Cretaceous Research 129 (2022) 105029

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Cretaceous Research
journal homepage: www.elsevier.com/locate/CretRes

The sauropod record of Salitral Ojo del Agua: An Upper Cretaceous

(Allen Formation) fossiliferous locality from northern Patagonia,
Argentina
Mauro Aranciaga Rolando a, b, *, Jordi A. García Marsa

a, Federico L. Agnolín a, b, c,
a n Rozadilla , Fernando E. Novas a
Matías J. Motta , Sebastia a

a
Museo Argentino de Ciencias Naturales ‘Bernardino Rivadavia’, Consejo Nacional de Investigaciones Científicas y T
ecnicas e CONICET, Buenos Aires,
Argentina
b
Museo Municipal de Ciencias Naturales “Carlos Ameghino”, Mercedes, Argentina
c n de Historia Natural “F
Fundacio elix de Azara”, Departamento de Ciencias Naturales y Antropología, Universidad Maimo nides, Buenos Aires, Argentina

a r t i c l e i n f o a b s t r a c t

Article history: The record of sauropods in the Upper Cretaceous of Patagonia is rich. However, there are still several
Received 2 October 2020 blanks on this record. Here it is described a new sauropod assemblage coming from the Salitral Ojo de
Received in revised form Agua Area, Río Negro province, northern Patagonia, Argentina (Allen Formation; Maastrichtian). Remains
21 August 2021
of indeterminate saltasaurines, aeolosaurines, as well as new specimens of the small saltasaurine
Accepted in revised form 5 September 2021
Rocasaurus muniozi and the new eutitanosaur Menucocelsior arriagadai gen. et sp. nov., are described. The
Available online 20 September 2021
new eutitanosaur is represented by an incomplete caudal series and some appendicular bones that
indicate that it does not belong to any previously recognized eutitanosaur clade (e.g, Colossosauria,
Keywords:
Titanosauria
Saltasaurinae, Aeolosaurini). The co-occurrence of several roughly coeval titanosaurs in a restricted area
Menucocelsior (as occur in Bajo de Santa Rosa, Salitral Moreno and Salitral Ojo de Agua sites) with different body plans,
Rocasaurus muniozi indicates that they probably occupied particular ecological niches and that probably competition for
Sauropod osteoderms resources was limited, allowing the connivance of several taxa in a single locality and age. This pattern is
Maastrichtian not observed in other sites of the world. Based on these records and on the high diversity of osteoderm
Patagonia morphotypes recovered, it appears to be certain that palaeoecological conditions in the area were able to
sustain and promote the flourishing of many species of titanosaurs.
© 2021 Elsevier Ltd. All rights reserved.

1. Introduction
The fossil record of sauropods in Patagonia is one of the richest
in the world, with more than 50 formally described species. Up-
permost Cretaceous (CampanianeMaastrichtian) beds of northern
Patagonia are well-known by its high diversity of sauropods, spe-
cifically titanosaurs (Salgado and Bonaparte, 2007). As an example
of such diversity, more than four different taxa are known from
roughly coeval outcrops of the Maastrichtian Allen Formation:
Rocasaurus muniozi (Salgado and Azpilicueta, 2000), Aeolosaurus sp.
(Salgado and Coria, 1993), Bonatitan reigi (Martinelli and Forasiepi,
2004), and Antarctosaurus wichmannianus (von Huene, 1929).
* Corresponding author. Museo Argentino de Ciencias Naturales ‘Bernardino
cnicas e CONICET,
Rivadavia’, Consejo Nacional de Investigaciones Científicas y Te
Buenos Aires, Argentina.
E-mail address: mauro.a_guido@hotmail.com (M.A. Rolando).
In this regard, during 1989 and 1994 several expeditions carried
out by the Museo de Geología y Paleontología de la Universidad
Nacional del Comahue (Neuque n Province, Argentina) and the
Museo Provincial “Carlos Ameghino” (Río Negro Province,
Argentina) resulted in the finding of one of the most prolific fossil
sites that yielded sauropod remains from the Allen Formation. This
site, known as Salitral Moreno, at northern Río Negro Province,
yielded six different titanosaur species, including Rocasaurus
muniozi, Aeolosaurus sp., and four still innominate taxa (García and
Salgado, 2013). This indicates that the diversity of sauropods in the
Allen Formation still awaits further discoveries.
The aim of the present contribution is to describe a new
sauropod fauna coming from the Allen Formation beds at “Salitral
Ojo de Agua” site, near Arriagada Farm, northeastern Río Negro
Province, Patagonia, Argentina. This fossil site has yielded remains
of pterosaurs and plesiosaurs, as well as diverse dinosaurs
including alvarezsaurid theropods, hadrosaurid ornithopods, and

https://doi.org/10.1016/j.cretres.2021.105029
0195-6671/© 2021 Elsevier Ltd. All rights reserved.

, F.L. Agnolín et al.
M.A. Rolando, J.A. García Marsa
sauropod nesting sites (Salgado et al., 2007, 2009; Novas et al.,
2012). However, no sauropod bones from this site were described
up to the date.
2. Materials and methods
The specimens here described come from the Cerro Matadero
site (67
230 22.1300 W; 39
300 27.0800 S; Fig.1), located within the
Arriagada Farm, at 70 km south from General Roca city, Río Negro
province, Argentina. This area is known as “Salitral Ojo de Agua”
(Agnolín et al., 2012) and exhibits outcrops pertaining to the Allen
Formation (Maastrichtian; Garrido 2010). This stratigraphic unit is
composed by sandstones, mudstones and pelites reaching about 70
m thick that represent shallow water courses, bearing freshwater
and brackish vertebrates and mollusks (Casadio, 1994; Martinelli
and Forasiepi, 2004).
Dinosaurs are well-known in Allen Formation, and include non-
avian theropods such as the abelisaurid Quilmesaurus curriei (Coria,
2001), indeterminate tetanurans (Coria and Salgado 2005), the
alvarezsaur Bonapartenykus ultimus (Agnolín et al., 2012), the large-
sized unenlagiid Austroraptor cabazai (Novas et al., 2009), as well as
the birds Limenavis patagonica (Clarke and Chiappe 2001) and
Lamarqueavis australis (Agnolín, 2010). Closer to Cerro Matadero
site, a single isolated and incomplete vertebra (specimen MPCN-
PV-802) of an indeterminate saltasaurine sauropod was found in
coeval beds from the nearly located Cerro Guerra site, which yiel-
ded remains of giant pterosaurs belonging to Azhdarchidae (Novas
et al., 2012).
For the nomenclature of vertebral laminae, it is used Wilson
(1999), and for vertebral fossae that employed by Wilson et al.
(2011). In the case of osteological nomenclature of caudal vertebrae
it is followed that of Campos et al. (2005). The taxonomic nomen-
clature follows Salgado and Bonaparte (2007). The name of the clades
such as Eutitanosauria, follow the topology and diagnosis of previous
authors (Carballido et al., 2017; Gonza lez-Riga et al., 2019).
Regarding osteoderms, several authors have proposed different
schemes for classifying titanosaur scutes (von Huene, 1929;
Bonaparte and Powell, 1980; Salgado, 2003). It is followed the system
proposed by D'Emic et al. (2009) because it includes the largest
known morphological osteoderm diversity. Following this classifica-
tion, four different morphotypes are recognized (Morphotypes 1 to 4).
Regarding the relative locations of specific structures within the
osteoderms, we adhere to the terms ‘superficial’ and ‘deep’ pro-
posed by Hill (2006, 2010). These terms are synonyms of ‘external/
basal’ (Scheyer and Sander, 2004), ‘distal/proximal’ (Main et al.,
2005), and ‘external/internal’ (D'Emic et al., 2009). It is used the
term ‘marginal region’ to refer to the area of contact between the
superficial and deep faces, as proposed by Cerda et al. (2015).
Because in most osteoderms cannot be established which portion is
cranial or caudal, we use ‘cranial/caudal’ (CR/CA) referring to one of
the edges of the osteoderm as proposed by Cerda et al. (2015).
According to the terminology employed by von Huene (1929) and
Salgado (2003), it is used the term ‘cingulum’ to refer to a periph-
eral ring of tubercles running along the external margins of the
osteoderm.
Institutional abbreviations. Museo Provincial “Carlos Ame-
ghino”; Cipolletti, Río Negro Province, Argentina, MPCA; Museo
Patago nico de Ciencias Naturales, Vertebrate Paleontology collec-
tion, General Roca, Río Negro Province, Argentina, MPCN-PV.
3. Systematic paleontology
Sauropoda Marsh, 1878
Titanosauria Bonaparte and Coria, 1993
2
Cretaceous Research 129 (2022) 105029
Eutitanosauria Sanz, Powell, Le Loeuff, Martínez and Pereda-
Suberbiola, 1999
Menucocelsior gen. nov.
Zoobank registration: urn:lsid:zoobank.org:pub: 7509E30A-DAA6-
4968-AD4C-103C63479EF1
Etymology. Menuco, from the Mapundungún, meaning waterhole.
This refers to the fossil site “Salitral Ojo de Agua”, and celsior, from
the Latin, meaning major.
Diagnosis. Mid-sized titanosaur of strongly procoelous caudal
vertebrae and stout humerus diagnosable on the following com-
bination of characters: 1) first two anterior caudal vertebrae with
transverse processes that are rod-like and dorsoventrally low
(probably autapomorphic); 2) first and second caudal vertebrae
with transverse processes that are subtriangular in contour and
laterally oriented in dorsal view; 3) second caudal with ante-
rodorsally oriented prezygapophyses; 4) mid-caudal vertebrae
with neural arches located posterior to the anterior margin of the
centrum; 5) humerus with well-developed supraectepicondylar
ridge, forming a prominent flange of bone (probably autapomor-
phic); 6) humerus with short deltopectoral crest that does not ex-
tends up to the mid-length of the bone; 7) short and stout fibula
with transversely narrow distal end; and 8) distal end of fibula with
proximodistally expanded and prominent anterior flange, repre-
senting more than one quarter of the total fibular length (probably
autapomorphic).
Type and only included species. Menucocelsior arriagadai sp. nov.
Menucocelsior arriagadai gen. et sp. nov.
Etymology. The specific epithet refers to “Beto” Arriagada and his
family, the owners of the Farm that includes the fossil sites here
reported, which are also enthusiastic on preserving fossils from the
area and usually help us to find fossil sites during our fieldtrips.
Holotype. MPCN-PV-798, associated specimen (Fig. 1B-C) consisting
on first, second and probably third caudal vertebrae, three distal
anterior-caudals, six proximal mid-caudals and five distal mid-
caudals, right humerus, left fibula and incomplete metapodial.
The position of the vertebrae is determined on the basis of com-
parisons with complete or nearly complete caudal sequences of
some titanosaurs as Trigonosaurus, Baurutitan, Overosaurus, Neu-
quensaurus Narambuenatitan and Notocolossus (Campos et al.,
2005; Kellner et al., 2006; Filippi et al., 2011; Coria et al., 2013;
lez-Riga et al., 2016). The elements identified as first and
Gonza
second caudals differ from other caudal elements, in that they show
anteroposteriorly short and dorsoventrally high centra, and later-
ally oriented transverse processes lacking a posterior tilting. The
element identified as the first caudal shows a centrum with a
transversely width that surpass its dorsoventral height; when
compared with other caudals (including the second one) it lacks a
well-defined ventral longitudinal groove, and shows prominent
spinodiapophyseal and centrodiapophyseal laminae (this later
lamina is retained in the second caudal but it is poorly developed).
More distal caudals show elongate and dorsoventrally low centra
with a well-defined ventral groove. Distal anterior caudals are
notably smaller than the anteriormost ones but still retain a short
and wide centrum, and a shallow ventral longitudinal excavation.
They show a centrum with the anterior articular surface that is
ovoidal in contour, with a transversely oriented main axis. Mid-
caudals show centra that are relatively elongate, lack a ventral
longitudinal groove and exhibit a reduced transverse processes. As
occurs in most titanosaurs mid-caudal centrum length progres-
sively increases from proximal to more distal elements.
Diagnosis. The same as for genus by monotypy.

, F.L. Agnolín et al.
M.A. Rolando, J.A. García Marsa
Description. All caudal vertebral centra (Fig. 2e4) are strongly
procoelous with anteriorly positioned neural arches, as occurs on
most titanosaurs. With exception of the first caudal, the remaining
elements show a longitudinally concave ventral surface, but lack
well-defined longitudinal groove and keels. There is no external
evidence of pneumaticity in any preserved caudal vertebra. The
centra lack internal camellate structure.
Most proximal caudals (Fig. 2) exhibit anteroposteriorly short and
dorsoventrally high centra, with notably concave lateral surfaces.
The concave anterior articular surface is delimited by a prominent
osseous ring. The first caudal centrum is transversely wider than
dorsoventrally tall, whereas the second caudal is taller than wide.
The anterior articular surface on proximal caudals is canted and
thus the dorsal margin extends anteriorly beyond the ventral one.
The apex of the convexity of the posterior articular surface is
slightly dorsally displaced above the mid-height of the centrum.
The lateral surface of the centra shows two symmetrically located,
ellipsoidal-shaped and relatively large-sized foramina.
Neural arches of first and second caudals are poorly preserved, and
the neural spine has been broken off. The preserved portion in-
dicates that the neural arches were anteriorly oriented, ante-
roposteriorly short, dorsoventrally low, and transversely wide. The
base of the neural spine suggests that it was very broad and thick,
subtriangular in cross-section with the apex posteriorly oriented.
Transverse processes are laterally oriented and subtriangular in
contour when viewed dorsally.
The first caudal (Fig. 2) shows a badly damaged neural arch. The
spinodiapophyseal lamina is prominent. The neural canal is trans-
versely wider than dorsoventrally low. The transverse processes are
posterolaterally oriented and sub-horizontally positioned. They are
rod-like and dorsoventrally low, especially at their base, and lack a
dorsal tubercle. The anteroventral ridge is well-developed and ex-
tends along the transverse process. The anterior cen-
trodiapophyseal lamina is represented by a poorly developed ridge
of bone. It shows a ventral longitudinal depression that is delimited
by two subparallel thick and rounded ridges that end on haemal
facets (Fig. 2D).
The second caudal (Fig. 3) is nearly completely preserved, and
shows eroded neural spine and the left postzygapophysis. The
neural arch is anteroposteriorly narrow and shows anteriorly ori-
ented anterior and posterior margins (Fig. 3E). Spinoprezygapo-
physeal laminae are narrow and prominent, and joined together
form a “V” in dorsal view (Fig. 3C). The prezygapophyses are rela-
tively elongate, narrow and rod-like, with small articular surfaces.
In dorsal view, both prezygapophyses are separated by a “U” shaped
intraprezygapophyseal space. Centroprezygapophyseal laminae are
columnar, relatively thick and well-defined (Fig. 3B). The spino-
diapophyseal lamina is poorly defined. As occurs with the first
caudal, the transverse processes are rod-like and dorsoventrally
low.
Distal anterior-caudals (Fig. 4A-J) are represented by isolated centra
(with fragments of the base of the neural arch). They show a more
prominent procoely than anteriormost caudals. In lateral view, the
centra are strongly anteroposteriorly concave, and show a notably
concave ventral surface (Fig. 4G). There is no ventral longitudinal
groove. The posterior articular condyle is deeply convex, separated
from the centrum by a neck and is notably transversely expanded
(Fig. 4B). The apex of the convexity of the posterior articular surface
is closer to the dorsal part of the centrum. The transverse processes
are represented by their bases, are relatively robust and ante-
roposteriorly extended. Together with the dorsal tuberosity they
form a prominent and laterally oriented bulge.
Proximal mid-caudals (Fig. 4KeO) are strongly procoelous and
anteroposteriorly elongate. In anterior and posterior views, the
centra are roughly sub-quadrangular in contour (Fig. 4MeN). They
3
Cretaceous Research 129 (2022) 105029
lack a ventral longitudinal ridge or groove. The lateral surface of the
centrum is less concave than in more proximal caudals. The apex of
the convexity of the posterior articular surface is just slightly
dorsally displaced above the mid-height of the centrum. The neural
arches are located at the anterior half of the vertebral centrum, but
do not reach its anterior margin. The prezygapophyses are rela-
tively short and slightly surpass the anterior margin of the centrum
(Fig. 4O). The transverse processes are represented by a conspicu-
ous dorsoventrally low and anteroposteriorly elongate ridge placed
at the limit between the centrum and the neural arch.
The neural spine is relatively thick but dorsoventrally low and
anteroposteriorly short (Fig. 4O). It shows a well-defined prespinal
lamina and a very wide and deep “V”-shaped prespinal fossa (Fig.
4K). The spinoprezygapophyseal laminae are very well-developed
and relatively thick, being separated from the prezygapophyses
by a longitudinal lateral groove.
Distal mid-caudals (Fig. 4P-T) are strongly procoelous and relatively
elongate, much more than other caudals. In anterior and posterior
views, the centra are roughly subcircular in contour (Fig. 4R-S).
They lack a ventral longitudinal ridge or groove. The lateral and
ventral surfaces of the centrum are notably anteroposteriorly
concave. The apex of the convexity of the posterior articular surface
is dorsally displaced above the mid-height of the centrum. The
neural arch is not anteriorly oriented, and does not reach the
anterior margin of the vertebral centrum (Fig. 4T). The neural spine
is relatively thick but dorsoventrally low. The prespinal fossa is very
deep, narrow and “V”-shaped (Fig. 4P).
The humerus (Fig. 5A-E) is represented by a right element lacking
the proximal articular surface and having a slightly eroded delto-
pectoral crest. It is relatively robust and straight-shafted. Its prox-
imal and distal ends are not greatly expanded (Fig. 5A-B). The
lateral margin of the shaft is nearly straight; whereas the medial
margin is smoothly concave (Fig. 5C-D). The shaft is strongly
anteroposteriorly compressed, transversely expanded and is ellip-
tical in cross-section.
In spite of being partially preserved, the shaft lacks strongly
divergent medial and lateral margins.The lateral margin of the
proximal end shows a relatively thick deltopectoral crest (Fig. 5A).
It is dorsoventrally short and does not reach the mid-length of the
bone. It lacks a distal torsion and expansion and lacks a distolateral
bulge (for the anchoring of M. latissimus dorsi; Otero, 2010) in
posterior view (Fig. 5B). The anterior fossa (coracobrachial fossa in
Gonza lez-Riga et al., 2016) is deep and wide, subtriangular-shaped
in contour, and with a distally oriented apex (Fig. 5A-E). It is not
delimited by well-defined ridges or crests. There are no well-
developed bumps for the abductor musculature.
The distal end of the bone is medially oriented. In posterior and
anterior views, the humerus lacks supracondylar ridges (Fig. 5E).
The supraectepicondylar ridge is represented by a prominent flange
of bone. The radial and ulnar condyles are slightly twisted with
respect to the proximal end of humerus. Both condyles are sub-
equal in size and shape, however, the ulnar one is slightly smaller
and more distally extended than the radial one. A well-defined
posterior distal fossa (olecranal fossa in Gonza lez-Riga et al.,
2016) is present.
The left fibula (Fig. 5F-I) shows a relatively stout and straight shaft
that is subtriangular in cross-section. Both proximal and distal ends
are poorly expanded (Fig. 5F-G). The proximal half of the bone is
slightly sigmoidal in outline. In lateral view, the proximal end of the
bone is gently convex, with exception of a pronounced and rela-
tively well-defined anterolateral depression. At the proximal third
of the shaft exist a flange-like, anteroposteriorly narrow, and single
lateral tuberosity (Fig. 5F, H), very different from the condition
observed in Laplatasaurus (Gallina and Otero, 2015). The distal half
of the shaft shows roughly subparallel anterior and posterior

, F.L. Agnolín et al.
M.A. Rolando, J.A. García Marsa Cretaceous Research 129 (2022) 105029

Fig. 1. A, map showing the fossiliferous localities where the fossil was extracted; B, map showing the quarry that yielded the holotype of Menucocelsior arriagadai (MPCN-PV-798;
shaded in light grey) found together with intermixed bones of hadrosaurs (shaded in dark grey); C, silhouette of Menucocelsior arriagadai showing the preserved bones in white.
4

, F.L. Agnolín et al.
M.A. Rolando, J.A. García Marsa
margins, and its lateral surface is homogeneously convex. The distal
end of the bone shows a prominent anterior flange that is prox-
imodistally expanded and represents more than one quarter of the
total fibular length (Fig. 5F, G, I).
In medial view (Fig. 5I), the proximal end shows a flat and sub-
triangular surface for articulation with the tibia. Distally, the shaft
shows a poorly defined longitudinal groove that ends at the distal
third of the bone. In lateral view, the distal end of the bone is not
expanded, being subequal in width to the fibular shaft; it is sub-
circular in cross-section.
In both anterior and posterior views (Fig. 5F-G), the fibular shaft is
narrow and shows slightly divergent proximal and distal ends. The
later lacks prominent anterior and posterior expansions.
Remarks. Menucocelsior is referred to the Eutitanosauria on the
basis of procoelous anterior and mid caudal vertebrae, and neural
arches of caudal vertebrae located at the anterior half (Salgado et
al., 1997; Bonaparte, 1999; Powell, 2003; Gonz alez-Riga et al.,
2019). Further, Menucocelsior differs from non-eutitanosaurs as
Narambuenatitan, Malawisaurus and Epachthosaurus in lacking
elongate and narrow neural spines on mid-caudal vertebrae (Jacobs
et al., 1993; Filippi et al., 2011).
Among eutitanosaurs, Menucocelsior differs from the giant colos-
sosaurians (such as Futalongkosaurus, Patagotitan or Notocolossus)
not only in body size, but also on several additional features,
namely relatively elongate caudal vertebral centra, humerus with
well-defined distal condyles and deltopectoral crest not distally
expanded and lacking a medial tilting (Gonz alez-Riga et al., 2016,
2019).
Menucocelsior differs from saltasaurines, such as Pellegrinisaurus,
Saltasaurus and Neuquensaurus, in lacking dorsoventrally depressed
caudal centra and large cells of spongy bone, humerus lacking a
prominent posterolateral bulge on deltopectoral crest, radial and
ulnar condyles not well-separated and lacking supracondylar
ridges (Powell, 1992; Salgado, 1996; D'Emic, 2012; Gonza lez-Riga et
al., 2019).
Aeolosaurines, in contrast to Menucocelsior, show unique caudal
vertebrae characterized by neural arches with notably narrow,
elongate and strongly anteriorly projected prezygapophyses and
neural spines, and caudal centra being dorsoventrally tall and
having a transversely narrow ventral surface (Salgado and Coria,
1993; Salgado et al., 1997; Kellner and Azevedo, 1999; Powell,
2003; Franco-Rosas et al., 2004; Casal et al., 2007; Calvo and
Porfiri, 2010; Santucci and Arruda-Campos, 2011; Martinelli et al.,
2011; Coria et al., 2013). The neural arches of Menucocelsior are
incompletely known. In spite to that, at the anterior caudals the
neural arches appear to be anteroposteriorly short and probably the
postzygapophyses were located at the anterior half of the vertebra,
a condition that occurs in aeolosaurines (Casal et al., 2007). How-
ever, in contrast to aeolosaurines, Menucocelsior anterior proximal
and mid-caudals show neural arches that are not continuous with
the anterior margin of the centrum (Casal et al., 2007), a condition
that is widespread among titanosaurs (Powell, 2003; Campos et al.,
2005; Santucci and Bertini, 2006).
Menucocelsior humerus differs from saltasaurines such as Sal-
tasaurus and Neuquensaurus in being more gracile, lacking
strongly expanded proximal and distal ends, in having delto-
pectoral crest that does not extend more than half the length of
the bone, distal condyles not anteriorly beveled, and in the
absence of well-defined posterior distal fossa and supracondylar
ridges (Powell, 2003; Otero, 2010). It also differs from material
referred to Laplatasaurus, Rapetosaurus, Bonitasaura, aeolosaur-
ines and its kin (as Petrobrasaurus, Gondwanatitan, Muyelensaurus
and Aeolosaurus) by having more robust proportions and a
distally extended radial condyle (Kellner and Azevedo, 1999;
Calvo et al., 2007; Curry-Rogers, 2009; García and Salgado, 2013;
5
Cretaceous Research 129 (2022) 105029
Gallina and Otero, 2015; Gallina and Apesteguía, 2015). Further,
the presence of a well-differentiated and proximodistally
extended supraectepicondylar ridge (sensu Powell, 1992), is a
feature uncommon in titanosaurs and present in few selected
taxa, such as Saltasaurus and Muyelensaurus (Powell, 1992; Calvo
et al., 2007).
The fibula of Menucocelsior is relatively short and stout, but not to
the degree observed in saltasaurines such as Neuquensaurus and
Saltasaurus (Powell, 1992; 2003). Further, it is nearly straight, in
contrast with the strongly sigmoidal proximal half exhibited by
saltasaurines (Gonza lez-Riga et al., 2019). The lateral tuberosity is
single and relatively prominent, being different from the double
condition observed in several titanosaurs (e.g., Epachthosaurus,
Laplatasaurus, Uberabatitan, Antarctosaurus; Salgado et al., 2015).
The distal end of the fibula in Menucocelsior in side view is poorly
anteroposteriorly expanded. On the other hand, in Aeolosaurus and
Neuquensaurus the distal end shows a notably expanded distal end
with a strongly convex and somewhat pulley-shaped distal surface
(von Huene, 1929; Powell, 2003).
The distal end of the fibula in Menucocelsior shows a prominent and
proximodistally extended bony flange that strongly diverges from
the shaft. A condition absent in most titanosaurs (e.g., Bonitasaura,
Laplatasaurus, Rapetosaurus, Epachthosaurus; Curry-Rogers, 2009;
Gallina and Otero, 2015). An extensive crest is also present in some
titanosaurs such as Neuquensaurus (Otero, 2010) and Mendoza-
saurus (Gonza lez-Riga et al., 2019), but in both cases the bone
flange is not as prominent as observed in Menucocelsior.
Finally, Menucocelsior differs from the roughly coeval Antarctosau-
rus wichmannianus in addition to the great size-disparity, in having
much more robust humerus, presence of supraectepicondylar crest
and in lacking a strong ventral ridge delimiting the coracobrachial
fossa (von Huene, 1929).
In addition to the above-mentioned combination of characters,
Menucocelsior exhibits several traits that are unique or scarce
among titanosaurs. As for example, the anterior caudal vertebrae of
Menucocelsior show a combination of characters that is not known
in other titanosaurs, including: centra with sub-quadrangular
anterior articular surfaces and ovoidal posterior articular surfaces
that are ventrally narrow; ventral surface excavated by a shallow
longitudinal concavity; transverse processes dorsoventrally low
and with a narrow base; transverse processes relatively elongate
and posterolaterally oriented, lacking a ventral tilting; first caudal
vertebra having a well-developed anteroventral ridge along all the
length of the transverse process. As indicated above, this combi-
nation of characters is absent in most members of saltasaurines,
aeolosaurines and colossosaurians (Powell, 1992, 2003; Martinelli
et al., 2011; Gonzalez-Riga et al., 2016, 2019), and other eutitano-
saurs as “nemegtosaurines” (e.g., Bonitasaura, Rapetosaurus; Curry-
Rogers, 2009; Gallina and Apesteguía, 2015), and the Brazilian
Trigonosaurus (Campos et al., 2005), and Baurutitan (Kellner et al.,
2006). Further, derived titanosaurs, including saltasaurines, Ala-
mosaurus, Antarctosaurus, and Baurutitan exhibit a derived
biconvex first caudal centrum (Calvo and Gonz alez-Riga, 2003;
Kellner et al., 2006; Novas, 2009; see different interpretation in
D'Emic and Wilson, 2011). In spite of the important differences
noted above, Menucocelsior resembles Trigonosaurus and Baurutitan
in that the first and second caudals show relatively narrow and
dorsoventrally low transverse processes, a condition uncommon
among titanosaurs (Campos et al., 2005; Kellner et al., 2006).
In sum, Menucocelsior is an eutitanosaur that is clearly distin-
guishable from coeval members of the clade. Its anatomy distin-
guishes it from main titanosaur subclades, and lacks most
apomorphic features of saltasaurines and aeolosaurines. Because of
the paucity of the available material and the absence of clear syn-
apomorphic features with other titanosaurs, the phylogenetic

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Fig. 2. First caudal vertebra of Menucocelsior arriagadai in posterior (A), anterior (B), dorsal (C), ventral (D) and lateral (E) views. Abbreviations. avr, anteroventral ridge; cdl,
centrodiapophyseal lamina; hf, haemal facet; nc, neural canal; ns, incomplete fragment of neural spine; prz, prezygapophysis; Spdl, Spinodiapophyseal lamina; tp, transverse
process; vf, vascular foramen; vg, ventral groove. Scale bar: 10 cm.

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position of Menucocelsior among eutitanosaurs should still be
regarded as uncertain.
Saltasauridae Powell, 1992
Aeolosaurini Franco-Rosas et al., 2004
Genus and species indeterminate
Referred material. MPCN-PV-799, four incomplete distal caudal
vertebrae (Fig. 6).
Description. Available vertebrae are poorly preserved, eroded and
lack zygapophyses (Fig. 6). The centrum is elongate and strongly
procoelous. The apex of the convexity of the posterior articular
surface is slightly dorsally displaced above the mid-height of the
centrum. The vertebrae lack internal camellate structure.
The lateral faces are sub-vertically oriented being sub-rectangular
in lateral view and nearly straight, being gently anteroposteriorly
concave (Fig. 6E). The lateral face of the centrum is continuous with
the lateral surface of the neural arch. The neural arch is located at
the anterior margin of the centrum; it is transversely narrow and
dorsoventrally tall, with relatively narrow and tall neural arch
pedicles (Fig. 6A, E). In spite of being poorly preserved the anterior
margin of the neural arch appears to be strongly forwardly tilted.
The ventral surface of centrum is transversely narrow and ante-
roposteriorly concave and lacks a well-defined longitudinal groove
(Fig. 6B).
Remarks. Distal caudal vertebrae (Fig. 6) here described are poorly
preserved, and thus lack several diagnostic features. However, a
combination of characters is shared with Aeolosaurus, including
transversely narrow vertebrae (especially the ventral surface of
centrum), lateral surface of centrum nearly straight and continuous
with lateral surface of neural arch, and neural arch located at the
anterior edge of the centrum with its anterior margin strongly
forwardly oriented (Powell, 1987, 2003; Franco-Rosas et al., 2004;
García and Salgado, 2013). Several species of Aeolosaurus are known
from roughly coeval beds in Argentina, Brazil and Tanzania (Powell,
1987; Salgado and Coria, 1993; Salgado et al., 1997; García and
Salgado, 2013; Gorscak et al., 2017), thus the finding of a member
of this genus or a closely related form in Arriagada Farm is not
surprising.
Saltasaurini Salgado and Bonaparte, 2007
Rocasaurus muniozi Salgado and Azpilicueta, 2000
Referred material. MPCN-PV-800, associated specimen consisting
on cervical 12 neural arch, indeterminate posterior cervical
centrum and neural arch, neural arch of the second dorsal vertebra
(Figs. 7e9) and a fragmentary ischium.
Description. The material here described consists on vertebral ele-
ments that belong to a very small titanosaur (Figs. 7-9). The neural
arches are not fused to the vertebral centra, a condition that probably
indicates that the specimens here described were not somatically
mature. It is worth to mention that most known specimens of
Rocasaurus lack strong vertebral neural arch-centra connection
(Salgado and Azpilicueta, 2000; García and Salgado, 2013).
The only available cervical centrum (Fig. 7B-G) is poorly preserved
and strongly abraded, showing a camellate internal structure. The
centrum is very elongate and relatively low (it is shorter in Sal-
tasaurus; Zurriaguz and Powell, 2015), strongly opisthocoelous and
with a lateral pleurocoel that occupies approximately half of its
total height (Fig. 7B). In anterior view (Fig. 7G), the centrum is
roughly sub-quadrangular in contour. The base of the parapophyses
have been preserved, they are relatively robust and are ven-
trolaterally projected. The neural canal is relatively large, being
transversely wider than dorsoventrally tall (Fig. 7D). It is nearly as
tall as half of centrum height.
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Cretaceous Research 129 (2022) 105029
The 12th cervical arch (Fig. 8) was identified based on comparisons
with available complete titanosaur vertebral series, specially sal-
tasaurids such as Saltasaurus and Neuquensaurus (Powell, 2003;
Salgado et al., 2005a; Zurriaguz and Powell, 2015). In any case, the
vertebral position is tentative because among titanosaurs cervical
number is variable (Curry Rogers, 2009). The element here
described resembles 12th cervical of other titanosaurs in that the
neural arch is relatively robust and anteroposteriorly short (indi-
cating that the vertebra was not very elongate), in having a thick
neural spine that is pyramidal-shaped and anteroposteriorly com-
pressed; the prezygapophyses are robust, short and strongly
separated from the midline of the vertebra; the diapophyses are
prominent and strongly laterally projected, with a wide articular
surface for the rib; and a relatively narrow prespinal fossa and a
wide and shallow postpinal fossa.
In anterior view (Fig. 8A-A'), the neural arch is roughly sub-
triangular in contour. The neural spine is subtriangular shaped, its
top being rounded and transversely thick. The spine along all it
dorsoventral extension shows slightly divergent lateral margins.
The spinoprezygapophyseal laminae are transversely wide and
gradually fuse with the neural spine. These are separated by a very
deep and wide spinoprezygapophyseal fossa. The median prespinal
lamina (sensu Salgado and Powell, 2010) is relatively narrow and
low, and does not reach the bottom of the prespinal fossa. The
intraprezygapophyseal lamina is sharp and sub-horizontally ori-
ented. The prezygapophyseal facets are relatively large, ovoidal in
contour and slightly medially oriented. Ventral to the pre-
zygapophyses it is observed the prezygodiapophyseal lamina,
which is nearly subvertically oriented and shows a laterally concave
margin. The diapophyses are anterolaterally oriented and do not
fuse to the cervical rib. The anterior centrodiapophyseal lamina is
ventrally concave and laterally projected. Anteriorly, a short,
strongly ventrally concave and notably laterodorsally projected
centroprezygapophyseal lamina is observed. The neural canal is
relatively small and ovoidal in contour, being delimited by dorso-
ventrally low and transversely thick peduncles formed by the
centroprezygapophyseal laminae.
In posterior view (Fig. 8B-B'), the neural arch is subtriangular in
shape. The neural spine is delimited by very thick and prominent
spinopostzygapophyseal laminae. The spinopostzygapophyseal
fossa is very wide, deep and subtriangular in contour. It shows a
nearly smooth surface. The intrapostzygapophyseal lamina is
relatively acute and posteriorly projected. The postzygapophyses
are subtriangular in contour and its articular facet is ventrally ori-
ented. Posteriorly, the postzygodiapophyseal lamina is relatively
narrow and obliquely oriented, forming the roof of a relatively
narrow but deep postzygapophyseal-centrodiapophyseal fossa. The
centropostzygapophyseal laminae are transversely narrow and
form the peduncles of the neural arch. They are slightly constricted
at mid-height. The neural canal is taller than wide, being much
narrower in anterior view.
In lateral view (Fig. 8C-C'), the neural arch is roughly subtriangular in
contour. The zygapophyseal peduncles are relatively short and
indicate that zygapophyses probably do not extend beyond anterior
and posterior edges of centrum. The prezygapophyses are notably
dorsally flexed. The neural spine is subtriangular in contour, being
anteroposteriorly narrow but transversely thick. On its lateral sur-
face, the spine shows a very wide, deeply excavated, well-defined
and smooth spinodiapophyseal fossa. This fossa is subdivided by a
short epipophyseal-prezygapophyseal lamina connecting the pre-
spinal and the postzygodiapophyseal laminae. The neural spine is
connected with the prezygapophyses by strong spinoprezygapo-
physeal laminae. These laminae are thin, acute and extend laterally.
Both delimit a huge and subtriangular spinoprezygapophyseal fossa.

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Fig. 3. Second caudal vertebra of Menucocelsior arriagadai in posterior (A), anterior (B), dorsal (C), ventral (D) and lateral (E) views. Abbreviations. avr, anteroventral ridge; cdl,
centrodiapophyseal lamina; hf, haemal facet; nc, neural canal; ns, incomplete fragment of neural spine; prz, prezygapophysis; Spdl, pinodiapophyseal lamina; Sprl, spinoprezy-
gapophyseal lamina; tp, transverse process; vf, vascular foramina; vg, ventral groove. Scale bar: 10 cm.

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Fig. 4. Distal proximal (AeJ), Mid-proximal (KeO) and Mid-distal (PeT) caudal vertebrae of Menucocelsior arriagadai, in dorsal (A, F, K and P), ventral (B, G, L and Q), anterior (C, H, M
and R), posterior (D, I, N and S) and lateral (E, J, O and T) views. Abbreviations. dtu, dorsal tuberosity; fos, fossa; hf, haemal facet; na, base of the neural arch; ns, neural spine; poz,
postzygapophysis; prz, prezygapophysis; Sprl, spinoprezygapophyseal lamina; Sprf, spinoprezygapophyseal fossa tp, transverse process; Scale bars: 10 cm.

Such fossa shows two thin and short prespinal lamina on its anterior
surface.
In dorsal view (Fig. 8D-D'), the prezygapophyses are anterolaterally
directed and show oval-shaped and flat articular facets. Between
both prezygapophyses it is observed a transversally wide, deep and
subtriangular-shaped spinoprezygapophyseal fossa. The dia-
pophysis is subtriangular in contour. The neural spine is broken, but
it seems to be transversely wider that anteroposteriorly long.
Posterior to the neural spine, two divergent and strong spino-
postzygapophyseal laminae are observed. These delimit a notably
deep and subtriangular-shaped spinopostzygapophyseal fossa. A
wide but shallow spinodiapophyseal fossa is present.
In ventral view (Fig. 8E-E'), the diapophysis shows a relatively large
articular surface. At its base it exhibits a relatively small but deep and
well-delimited centrodiapophyseal fossa. This fossa is delimited ante-
riorly by the anterior centrodiapophyseal lamina. More anteriorly, a
transversely wide prezygapophyseal-centrodiapophyseal fossa is
observed. The postzygapophyses are transversely narrow and ellip-
soidal in contour. Both are joined by a posteriorly concave intra-
postzygapophyseal lamina. The intraprezygapophyseal lamina is nearly
straight.
The second dorsal vertebra (Fig. 9) is represented by a nearly
complete neural arch only lacking the tips of the neural spine and
the diapophyses. The neural arch is subtriangular shaped in ante-
rior and posterior views. This element is identified as a second
dorsal because the parapophyses are located at the base of the
neural arch, the neural arch is notably anteroposteriorly com-
pressed, the diapophyses shows a subtriangular and wide cross-
section, and the neural spine is not strongly posteriorly tilted.
In anterior view (Fig. 8A-A'), the neural spine is subtriangular in
contour and shows a nearly subvertically oriented anterior surface. It
shows a very transversely wide and deep prespinal fossa. The

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Fig. 5. Appendicular bones of Menucocelsior arriagadai. Right humerus in anterior (A), posterior (B), right lateral (C), left lateral (D) and distal (E) views. Left fibula in anterior (F),
posterior (G), lateral (H) and medial (I) views. Abbreviations. afl, anterior flange; afo, anterior fossa; ast, articular surface for tibia; dpc, deltopectoral crest; igr, intercondylar groove;
lt, lateral tuberosity; pdf, posterior distal fossa; rc, radial condyle; sec; supra ectepicondylar crest; uc, ulnar condyle. Scale bars: 10 cm.

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Fig. 6. Isolated caudal vertebra of an indeterminate Aeolosaurini (MPCN-PV-799), in dorsal (A), ventral (B), anterior (C), posterior (D) and lateral (E) views; and the Saltasaurini
indet. (MPCN-PV-801). Scale bars: 10 cm. Abbreviations. ce, centrum; na, neural arch; nc, neural canal.

spinodiapophyseal lamina is sharp, dorsolaterally concave and later-
ally defines the dorsal edge of the diapophysis. The prespinal lamina is
well-developed, dorsoventrally extended and reach the base of the
spinoprezygapophyseal fossa as the most preserved point of the
neural spine. The spinoprezygapophyseal laminae show a concave
lateral margin, and probably extended up to the top of the neural
spine. Together the spinoprezygapophyseal laminae conforms a
roughly hourglass contour in anterior view and reach the pre-
zygapophyseal facets. Such facets are anteromedially oriented. Both
are connected through a sub-horizontally oriented and anteriorly
projected intraprezygapophyseal lamina forming a prominent shelf
that conform the base of the spinoprezygapophyseal fossa. Just lateral

Fig. 7. Interpretative figure showing the distribution of the bones at the quarry that yielded the new specimen of Rocasaurus muniozi (MPCN-PV-800; shaded in black), and an
indeterminate Saltasaurini (MPCN-PV-801; shaded in grey) (A). Incomplete cervical vertebra of Rocasaurus muniozi in dorsal (B), right lateral (C), anterior (D), ventral (E), left lateral
(F) and posterior (G) views. Abbreviations. na, neural arch; nc, neural canal; par, parapophysis; pl, pleurocoel; vs, ventral surface. Scale bar: A, 70 cm; B-G, 5 cm.

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Fig. 8. Neural arch of cervical 12 of Rocasaurus muniozi, in anterior (A, A'), posterior (B, B'), lateral (C, C'), dorsal (D, D') and ventral (E, E') views. Light grey parts indicate deep
structures. Dark grey areas indicate broken or missing parts. Scale bar: 10 cm: Abbreviations: Acdl, anterior centrodiapophyseal lamina; Cdf, centrodiapophyseal fossa; Cpol,
centropostzygapophyseal lamina; Cprl, centroprezygapophyseal lamina; di, diapophysis; Eprl, epipophyseal-prezygapophyseal lamina; nc, neural canal; ns, neural spine; Podl,
postzygodiapophyseal lamina; Po-cdf, postzygapophyseal-centrodiapophyseal fossa; poz, postzygapophyses; Prdl, prezygodiapophyseal lamina; Prsl, prespinal lamina; prz,
prezygapophsyes; Spdf; spinodiapophyseal fossa; Spof, spinopostzygapophyseal fossa; Spol, spinopostzygapophyseal lamina; Sprf, spinoprezygapophyseal fossa; Sprl, spinopre-
zygapophyseal lamina; Tpol, intrapostzygapophyseal lamina; Tprl, intraprezygapophyseal lamina.

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to the prezygapophyseal articular facets a short prezygodiapophyseal
lamina is present. In anterior aspect, this lamina is laterally concave.
The centroprezygapophyseal laminae are curved and ventraally
concave, dorsoventrally short and placed at mid-length in the neural
peduncles. This lamina laterally delimits a shallow and subcircular-
shaped centroprezygapophyseal fossa. The neural arch peduncles
show a strongly concave lateral surface. The neural canal is subcircular
in contour.
In posterior view (Fig. 8B-B'), the diapophyses are subhorizontally
oriented and show gently concave dorsal and ventral margins. The
postzygapophyseal-centrodiapophyseal fossa is shallow, and ex-
hibits on its medial corner a deep and suboval-shaped pneumatic
foramen. The postzygapophyseal shelf is poorly preserved and
abraded. The spinopostzygapophyseal lamina is relatively promi-
nent and dorsoventrally short. The postzygapophyses are ovoidal
in contour and posterolaterally oriented. They are distally
extended by means of a poorly defined centropostzygapophyseal
lamina. This lamina is thin, laterally projected and dorsoventrally
short.
The postzygapophyses form the dorsal limit of a deep and well-
defined spinodiapophyseal-centrodiapophyseal pneumatic fossa.
There exists an additional subcircular pneumatic fossa located
dorsomedial to the postzygapophysis and that delimited a poorly
defined lamina that extended towards the top of the neural spine.
The presence of a pneumatic fossa cannot be observed on the right
half of the vertebra due to poor preservation. Distomedially to the
postzygapophysis, there is another subcircular-shaped pneumatic
fossa.
In dorsal view (Fig. 8C-C'), the prezygapophyses are notably ante-
roposteriorly compressed, being elliptical-shaped in contour. The
diapophyses are stout, transversely short and slightly anteriorly
oriented. Dorsolateral to the diapophysis a subcircular pneumatic
foramen is present.
In lateral view (Fig. 8D-D), the neural arch is strongly ante-
roposteriorly compressed. The neural spine is notably ante-
roposteriorly compressed and is slightly posterodorsally oriented.
The prezygapophyses are poorly anteriorly projected and sub-
horizontally oriented. The diapophyses are subtriangular in cross-
section. The postzygapophyses, diapophyses and pre-
zygapophyses are roughly at the same level. At the right side of the
neural arch the anterior centroprezygapophyseal lamina is notably
thick, rounded and conforms a strong buttress. The cen-
trodiapophyseal lamina is strongly posteriorly concave. It ends at
the dorsal margin of the parapophysis, which is partially included
within the base of the neural arch. At the left lateral side of the
neural arch shows anterior and posterior centrodiapophyseal
laminae. Both diverge ventrally: the anterior reaches the para-
pophyses and the posterior one reaches the posterior corner of the
neurocentral suture. Within these laminae, is observed a deep,
well-defined, and ellipsoidal-shaped centrodiapophyseal fossa. On
the right side of the element, this fossa is absent and the unique
centroprezygapophyseal lamina joins the base of the cen-
trodiapophyseal lamina. These laminae delimit dorsally a deep and
well-defined prezygapophyseal-centrodiapophyseal fossa.
Remarks. Specimens here described constitute an important addi-
tion to the knowledge of Rocasaurus muniozi. This taxon was orig-
inally described by Salgado and Azpilicueta (2000) who based their
description on a partially preserved skeleton including part of the
vertebral column, as well as several referred specimens. Later,
García and Salgado (2013) added some additional material to
Rocasaurus and recognized that several bones that were originally
referred to the taxon belong to other titanosaur taxa.
Present contribution adds two vertebral positions that were pre-
viously unknown in Rocasaurus. New elements are very well-
preserved and exhibit several features that may be diagnostic for
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Cretaceous Research 129 (2022) 105029
this taxon, when compared with other members of Saltasaurini
such as Saltasaurus and Neuquensaurus. In this sense, Salgado and
Azpilicueta (2000) noted some features for the holotype spec-
imen Rocasaurus that are not mentioned in other saltasaurines and
may be regarded as diagnostic for Rocasaurus. These features
together with others here recognized, and the fact that the Roca-
saurus holotype comes from the same geological unit, allow refer-
ring the specimens here described to this taxon.
Salgado and Azpilicueta (2000) proposed the presence of ante-
roposteriorly compressed neural spine on dorsal vertebrae as
diagnostic of Rocasaurus. In the specimen here described, as occurs
in the holotype, the anterior and posterior margins of the top of the
neural spine are roughly subparallel to each other for almost a
quarter of its total dorsoventral height. On the contrary, in Sal-
tasaurus and Neuquensaurus the neural spines of the last cervicals
are anteroposteriorly thicker than in Rocasaurus, showing anterior
and posterior margins in side view that are strongly divergent for
almost all their length, and the top of the neural spine is notably
bulkier (see Powell, 1992, 2003; Salgado et al., 2005a). This con-
dition appears to the even more evident in Saltasaurus, in which the
neural spine is notably low and mound-like (Powell, 1992).
As noted by Salgado and Azpilicueta (2000) in the dorsals of the
holotype of Rocasaurus, as well as in the specimens here described,
anterior and posterior centrodiapophyseal laminae are represented
by a single, robust and prominent bone buttress. This contrast with
the deeply bifurcate centrodiapophyseal lamina observed in dorsals
of Saltasaurus and Neuquensaurus (Powell, 1992; Salgado et al.
2005a; Zurriaguz and Powell, 2015).
In addition, there are several features shared by specimens here
described and Rocasaurus holotype that sustain they belong to the
same taxon. In Rocasaurus specimens the postzygapophyses of the
cervical vertebrae are not strongly projected outwards and back-
wards as in other saltasaurines such as Neuquensaurus and Sal-
tasaurus, being similar to the plesiomorphic titanosaur condition
(see Salgado et al., 1997).
Rocasaurus specimens resembles Neuquensaurus but differ from
Saltasaurus in several features, including cervical vertebrae with
straight intraprezygapophyseal lamina on cervical vertebrae (U-
shaped in Saltasaurus; Zurriaguz and Powell, 2015) and a shallow
centroprezygapophyseal fossa (deep and well-defined in Sal-
tasaurus; Zurriaguz and Powell, 2015). However, Neuquensaurus
shows on cervical 12 stout diapophyses that are deep, strongly
projected outwards, and slightly bent downwards (Salgado et al.,
2005a). In Rocasaurus, as is the case of Saltasaurus and other tita-
nosaurs (e.g., Isisaurus; Jain and Bandyopadhyay, 1997) the dia-
pophyses are notably weak and poorly laterally projected.
Further, the new specimen provides additional features that prob-
ably constitute autapomorphies of Rocasaurus. These include:
- Cervical 12 in anterior view with the spinoprezygapophyseal
laminae strongly separated from each other. In the case of
derived titanosaurs, as occurs in Neuquensaurus, both spinopre-
zygapophyseal and prespinal laminae form a prominent complex
structure that extends along all posterior cervical and dorsal se-
ries (Salgado et al., 2005a). In cervical 12 of Neuquensaurus this
complex is transversely narrow because the spinoprezygapo-
physeal laminae are very close to each other and are notably
curved towards the midline of the neural arch (Salgado et al.,
2005a). The same occurs in the anterior surface of the short-
ened neural spines known in Saltasaurus (Powell, 1992). In the
case of cervical 12 of Rocasaurus both spinoprezygapophyseal
laminae are far to each other and are nearly subvertically ori-
ented, being just gently curved towards the midline. In this sense,
the separation between both laminae is 46 mm on dorsalmost
point and 54 mm closer to its ventral margin. This results in a

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relatively wide and flat complex surface that is not present in
other saltasaurines.
- Second dorsal with notably anteroposteriorly compressed
articular facets of prezygapophyses. The second available dorsal
vertebra of Rocasaurus exhibits anteroposteriorly compressed
and transversely expanded prezygapophyseal articular surfaces,
which are ellipsoidal in contour and have a transversely oriented
major axis. This condition appears to be correlated to the strong
anteroposterior compression of the entire neural arch. In
contrast, in Saltasaurus and Neuquensaurus, as occurs in other
titanosaurs, the articular facet of the prezygapophyses in second
dorsal (and in other anterior dorsal vertebrae) is subcircular or
slightly ovoidal in contour, having its transverse and ante-
roposterior axes roughly subequal in size (Powell, 1992; 2003).
- Complex posterior surface of second dorsal, with several
pneumatic cavities and laminae. The posterior surface of the
dorsal vertebrae in titanosaurs is usually complex (e.g.,
Bonaparte, 1999). In the case of Rocasaurus it is particularly
pneumatic and has been considered as “hyperpneumatic” (Cerda
et al., 2012). At the second dorsal vertebra, Rocasaurus exhibits a
conformation that as a whole is not present in any other titano-
saur. At least, four pneumatic foramina have been identified.
Medial to the spinopostzygapophyseal is an additional lamina
that is laterally delimited at its base by a deep and well-defined
pneumatic foramen. Ventral to the postzygapophyses, the cen-
tropostzygapophyseal laminae are very prominent but dorso-
ventrally low and delimit very deep and ellipsoidal-shaped
postzygapophyseal-centrodiapophyseal pneumatic fossae that
are clearly visible when the neural arch is observed from the
sides. Medially there is an additional lamina that delimits a well-
developed and subcircular-shaped foramen. This combination of
characters resembles is some way the morphology observed in
other titanosaurs such as Neuquensaurus (Powell, 2003) and
Rapetosaurus (Curry-Rogers, 2009) but in both cases the surface is
far less complex and lacks prominent fossae and laminae, con-
trasting with the condition of Rocasaurus.
In sum, dorsal and cervical vertebrae of Rocasaurus muniozi here
described clearly depart from that known in other titanosaurs,
including the saltasaurines Saltasaurus and Neuquensaurus.
Genus and species indeterminate
Referred material. MPCN-PV-801, partially preserved fourth dorsal
vertebra (Fig. 10) and one incomplete and abraded cervical centrum
coming from the same quarry that yielded the materials of Roca-
saurus described above; MPCN-PV-802, incomplete ninth dorsal
vertebra (Fig. 11).
Description. Dorsal vertebrae are represented by an incomplete
anterior element (fourth dorsal) and an incomplete posterior
element (ninth dorsal). The first was found associated with the
materials of Rocasaurus but represents a much bigger individual.
Because of size differences and absence of apomorphic Rocasaurus
features, this element is referred as an indeterminate saltasaurine.
The second element was found in Cerro Guerra locality (approxi-
mately five kilometers away from Cerro Matadero locality). Both
elements lack hyposphene-hypantrum accessory articulations as in
derived titanosaurs and share notably anteroposteriorly shortened
Fig. 9. Neural arch of dorsal 2 of Rocasaurus muniozi, in anterior (A, A'), posterior (B, B'), dorsal (C, C') and lateral (D, D') views. Light grey parts indicate deep structures. Dark grey
areas indicate broken or missing parts. Scale bar: 10 cm: Abbreviations: Cdf, centrodiapophyseal fossa; Cdl, Centrodiapophyseal lamina; Cpol, centropostzygapophyseal lamina;
Cprf, centroprezygapophyseal fossa; Cprl, centroprezygapophyseal lamina; di, diapophysis; for, foramina; nc, neural canal; ns, neural spine; par, parapophysis; Pcdl, posterior
centrodiapophyseal lamina; Podl, Postzygodiapophyseal lamina; poz, postzygapophyses; Prdl, prezygodiapophyseal lamina; Prsl, prespinal lamina; Pr-cdf, prezygapophyseal-
centrodiapophyseal fossa; prz, prezygapophsyes; Spdl, spinodiapophyseal lamina; Spol, spinopostzygapophyseal lamina; Sprf, spinoprezygapophyseal fossae; Sprl, spinoprezy-
gapophyseal lamina; Tprl, intraprezygapophyseal lamina.
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Cretaceous Research 129 (2022) 105029
centra (which are relatively small when compared with neural arch
size) and very small ovoidal-shaped lateral pleurocoels that lack
any subdivision. They show internal camellate tissue. Based on this
shared combination of characters the specimens are tentatively
thought to belong to the same innominate taxon.
The anterior dorsal vertebra (MPCN-PV-801; Fig. 10) is represented
by a nearly complete neural arch lacking neural spine and right
diapophyses; the vertebral centrum is strongly abraded. The pre-
served portion of the centrum indicates that it was relatively small
and opisthocoelous, as occurs in most titanosauriforms (Salgado et
al., 1997). On its lateral surface, near to the dorsal margin is located
a small subtriangular pleurocoel, just below the base of the neural
arch. This vertebra is identified as a fourth element by having the
following combination of characters parapophyses near to the mid-
height of the neural arch, parapophyses robust, prominent and
dorsoventrally higher than anteroposteriorly long, weak para-
podiapophyseal laminae, and a wide, deep and subtriangular-
shaped prezygapophyseal-centrodiapophyseal fossa. This results
from comparisons with complete vertebral columns of derived
titanosaurs such as Overosaurus, Saltasaurus, and Neuquensaurus
(Powell, 2003; Salgado et al., 2005a; Coria et al., 2013; Zurriaguz
and Powell, 2015)
In anterior view (Fig. 10A-A'), the neural arch shows diapophyses
that are robust and strongly dorsally oriented, which results in
relatively transversely narrow vertebra. The diapophyses show a
well-defined and sub-parallel prezygoparapophyseal and
parapodiapophyseal laminae that delimit the anterior surface from
the lateral faces of the diapophyses. These two delimits a small and
oval prezygapophyseal-paradiapophyseal fossa. The pre-
zygapophyses are strongly transversely expanded and ovoidal in
contour. They are connected to each other by means of a strong and
dorsally concave intraprezygapophyseal lamina. This lamina is
strongly separated from the dorsal edge of the neural canal. The
intraprezygapophyseal lamina is dorsally bounded by a very wide
and deep spinoprezygapophyseal fossa. Immediately below to the
base of the prezygapophyses, there is a dorsoventrally short cen-
troprezygapophyseal lamina that reaches ventrally the dorsal
margin of the neural canal. Laterally, these laminae are delimited by
very large pneumatic centroprezygapophyseal fossae that are
subdivided by a strut of bone. Both centroprezygapophyseal
laminae together with the intraprezygapophyseal lamina form a
prominent complex that probably functioned like an accessory
articulation, probably similar to a hyposphene-hypantrum (see
Apesteguía, 2005). The neural arch peduncles are thick and
dorsoventrally tall and delimit an ovoidal neural canal that is
dorsoventrally taller than transversely wide.
In posterior view (Fig. 10B-B'), the surface of the neural arch is almost
flat, and the postzygapophyses do not project much posteriorly. The
posterior centrodiapophyseal lamina is thick and well defined and
delimits the lateral edges of the neural arch. The postzygapophyses are
not preserved but the centropostzygapophyseal laminae are observed.
They are nearly straight and prominent. They are laterally bounded by
wide and relatively shallow, but well defined, subcircular fossae. Be-
tween both centropostzygapophyseal laminae form a prominent
structure that probably articulated with the complex formed by the
centroprezygapophyseal and intraprezygapophyseal laminae.
In lateral view (Fig. 10C-C'), the centrum is dorsoventrally low and
shows a deep and subrectangular-shaped pleurocoel. A short,

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Fig. 10. Anterior dorsal vertebra of an indeterminate Saltasaurini (MPCN-PV-801) in anterior (A, A'), posterior (B, B'), left lateral (C, C') and right lateral (D, D') views. Light grey parts
indicate deep structures. Dark grey areas indicate broken or missing parts. Scale bar: 10 cm. Abbreviations: Acdl, anterior centrodiapophyseal lamina; Cpar, centroparapophyseal
lamina; Cpol, centropostzygapophyseal lamina; Cprf, centroprezygapophyseal fossa; Cprl, centroprezygapophyseal lamina; di, diapophysis; nc, neural canal; par, parapophysis;

16

, F.L. Agnolín et al.
M.A. Rolando, J.A. García Marsa
vertical and stout anterior centroparapophyseal lamina is present.
The neural arch is anteroposteriorly short and dorsoventrally tall.
The parapophyses are prominent and located at base of the dia-
pophyses. They are ovoidal in contour and very high with a
dorsoventrally oriented main axis. Posteroventral to the para-
pophysis there is a shallow parapophyseal-centrodiapophyseal
fossa. Within this fossa there are two small cavities that are sepa-
rated by an obliquely oriented strut of bone: a posterior and blind
one and an anterior, bigger and pneumatic one. From the ante-
rodorsal corner of the parapophyses a short but well-defined pre-
zygoparapophyseal lamina is present. This lamina shows on its
posterior margin a small but deep (the fossa is much shallower and
less defined in the left side of the vertebra). Such fossa is the
prezygapophyseal-paradiapophyseal fossa which is deep and pos-
teriorly delimited by a narrow but well-defined and prominent
paradiapophyseal lamina. The diapophyses are thick and robust,
roughly ovoidal in cross-section. Posteriorly, the diapophyses show
a thick, subtriangular and prominent posterior centrodiapophyseal
lamina, that forms the lateral margin of the posterior neural arch
pedicles. This lamina and the paradiapophyseal lamina delimit the
prezygapophyseal-centrodiapophyseal fossa, which is sub-
triangular and deep.
Posterior dorsal vertebrae are represented by an incomplete dorsal
9 that lacks the dorsal half of the neural arch and the left post-
zygapophyses (MPCN-PV-802; Fig. 11). It shares with the ninth
dorsal of derived titanosaurs as Neuquensaurus and Overosaurus
(Salgado et al., 2005a; Coria et al., 2013) strongly dorsally placed
parapophyses and diapophyses, transversely wide peduncles of
neural arch, parapophyseal-centrodiapophyseal fossa that is
notably complex, subtriangular shaped in contour and with an
anterodorsally oriented tip and prezygapophyses that are placed far
from each other in anterior view.
The neural spine was not preserved. The centrum is notably short
and shows a longitudinally concave ventral surface. There is no sign
of longitudinal groove or keel. In lateral view, the pleurocoel of the
centrum is relatively small (representing approximately less than
one fourth of total centrum length), shows an anteroposteriorly
oriented main axis and is dorsally delimited by a prominent
osseous edge.
In anterior view (Fig. 11A-A'), the diapophyses are robust and
strongly dorsolaterally oriented. The prezygapophyseal facets are
flat, strongly medially facing and round. Lateroventrally to the
prezygapophyses, it is observed a smooth, thin and curved pre-
zygoparapophyseal lamina. Between this lamina and the cen-
troprezygapophyseal lamina are observed, in the right side of the
element, two deep and oval pneumatic foramina within a shallow
fossa. This fossa is the prezygapophyseal-paradiapophyseal fossa.
On the left side, only one bigger pleurocoel is observed. The
intraprezygapophyseal lamina is thin, anterior projected and “V”
shaped. The anterior surface that surrounds the neural canal is flat
or slightly concave. The neural canal is round.
In posterior view (Fig. 11B-B'), the posterior surface of the neural
arch is almost flat or slightly concave. The posterior cen-
trodiapophyseal lamina is robust, slightly posteriorly projected and
laterally concave. The neural canal is oval, being wider than tall. The
centrum is rounded in contour and deeply concave, being deeper in
the center.
In dorsal view (Fig. 11E-E'), the bone is badly damaged. The pre-
zygapophyses are flat and round in contour. A deep prespinal fossa
is observed. Within this fossa it is observed a thin median
Pcdl, posterior centrodiapophyseal lamina; pl, pleurocoel; poz, postzygapophyses; Pp-cdf, parapophyseal-centrodiapophyseal fossa; Ppdl, paradiapophyseal lamina; Prdl, pre-
zygodiapophyseal lamina; Prpl, prezygoparapophyseal lamina; Pr-cdf, prezygapophyseal-centrodiapophyseal fossa; Pr-ppdf, prezygapophyseal-parapodiapophyseal fossa; prz,
prezygapophsyes; Tprl, intraprezygapophyseal lamina; Sprf, Spinoprezygapophyseal fossa.
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Cretaceous Research 129 (2022) 105029
prespinal lamina. In lateral view, the neural arch is ante-
roposteriorly narrow and dorsoventrally tall. The pre-
zygapophyses are prominent and show a convex anterior edge.
The parapophyses and diapophyses are not preserved. The pos-
terior centrodiapophyseal lamina is relatively thick and shows a
bifurcation near its base that results in a well-defined unnamed
parapophyseal lamina (sensu Salgado et al., 2005a) as occurs in
Neuquensaurus. This lamina delimits ventrally a wide
parapophyseal-centrodiapophyseal fossa that is subdivided by
several struts of bones, resulting in a notably complex neural arch.
The unnamed parapophyseal lamina delimits dorsally a wide and
deep prezygapophyseal-centrodiapophyseal fossa. The anterior
margin of this fossa is delimited by a notably thickened but
dorsoventrally low centrodiapophyseal lamina. There are two
relatively small pneumatic cavities lateral to the prezygapophyses.
In the left lateral view, the neural arch lacks such complex lami-
nation and fossae. A rudimentary unnamed parapophyseal lamina
is represented just by its base.
An isolated fragment of neural arch of an indeterminated vertebra
includes relatively small and subcircular-shaped prezygapophyses.
The prezygapophyseal facets are horizontal. Immediately lateral, it
is observed a deep and round foramen. A thin, laterally concave and
anterolaterally projected unnominated lamina it is observed.
Remarks. Specimens here described are referred to Saltasaurini
based on the presence of complex fossae below the transverse
process (with multiple crisscrossing laminae a morphology shared
with Rocasaurus, Neuquensaurus, Saltasaurus), and presence of an
unnamed additional parapophyseal laminae (as present in Roca-
saurus, Neuquensaurus, Saltasaurus) (Salgado et al., 2005a;
Zurriaguz and Powell, 2015) (Fig. 11F-J).
In the specimens here described, the pleurocoels of the vertebral
centra are relatively small to moderate in size and lack internal
subdivisions, features that clearly distinguish them from several
derived titanosaurs as Neuquensaurus, Rapetosaurus and Muye-
lensaurus (Salgado et al., 2005a; Calvo et al., 2007; Curry-Rogers,
2009), being similar in this aspect to some colosossaurians as
Mendozasaurus (Gonz alez-Riga, 2003). In anterior and posterior
views, the dorsal 4 MPCN-PV-801 exhibits a wide space between
the articular surfaces of the prezygapophyses. In anterior view, this
space is occupied by a subtriangular, flat and anteriorly facing
surface which is delimited by the intraprezygapophyseal and cen-
troprezygapophyseal laminae. This surface has its counterpart on
the posterior side of the bone. In this sense, the posterior side of the
vertebrae shows a subtriangular, flat and posteriorly facing surface.
As in the anterior side, here this surface is delimited by laminae, in
this case, the centropostzygapophyseal laminae. These anterior and
posterior flat surfaces show deep cavities on each lateral side. This
morphology contrast with some titanosaurs which shows a very
reduced space between the both zygapophyses and dorsal margin
of neural arch (e.g., Trigonosaurus, Rinconsaurus, Notocolossus,
Bonitasaura; Calvo and Gonz alez-Riga, 2003; Campos et al., 2005;
Gallina and Apesteguía, 2015; Gonza lez-Riga et al., 2016) or show a
wide space but devoid of strong and acute centrozygapophyseal
laminae and associated concavities (e.g., Aeolosaurus, Muyelensau-
rus, Barrosasaurus, Neuquensaurus, Rapetosaurus; Calvo et al., 2007;
Curry-Rogers, 2009; Salgado and Coria, 2009; Santucci and Arruda-
Campos, 2011). Mendozasaurus shows a similar conformation of the
anterior complex of laminae, but in posterior view, it exhibits wide
and well-defined infrapostzygapophyseal fossae (Gonza lez-Riga,
2003). Posterior dorsal vertebra here described shows a well-

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, F.L. Agnolín et al.
M.A. Rolando, J.A. García Marsa
defined unnamed additional parapophyseal lamina (Fig. 11F-J), a
feature previously reported only for the Saltasaurini sauropods
Rocasaurus, Saltasaurus and Neuquensaurus (Zurriaguz and Powell,
2015).
The incomplete preservation and dissociated nature of specimens
MPCN-PV-801 and MPCN-PV-802, together with the combination
of characters that distinguish both from other saltasaurines, pre-
cludes identification beyond Saltasaurini indet.
Osteoderms. Osteoderms are described separately from other oste-
ological remains because they have their own nomenclature and
classification. All the osteoderms here described come from the
Cerro Matadero site and were found isolated, far from any titano-
saur skeleton. Based on osteoderm classification employed D’Emic
et al. (2009), two different morphotypes have been found. Based on
D’Emic et al. (2009) these osteoderms may be characterized by:
Morphotype 1 (Ellipsoid). These osteoderms are relatively large
(from 3.7 to 59 cm) and oval-shaped in external and internal views.
Ellipsoid osteoderms are known from South America, Africa,
Madagascar, Europe, and India, and have been attributed to Men-
dozasaurus, Maxakalisaurus, Magyarosaurus, Aeolosaurus, Rapeto-
saurus, and Malawisaurus, and Saltasaurus (Le Loeuff et al., 1994;
Csiki, 1999; O’Leary et al., 2004; D’Emic et al., 2009; Vidal et al.
2014; Cerda et al., 2015).
Morphotype 2 (Keeled). This includes subcircular to oval-shaped
scutes (ranging from 7.2 to 20 cm) with one or more longitudinal
inner keels. They have been found from Argentina and Madagascar
and have been attributed to Neuquensaurus, Saltasaurus, and Aeo-
losaurus (Powell, 1992, 2003; Salgado et al., 2005a).
Description. MPCN-PV-803 corresponds to the Morphotype 2 of
D’Emic et al. (2009). It is a flattened and keeled element with an oval
outline in superficial view (Fig. 12 A-D). The cingulum is well
developed, and exhibits thick tuberosities. The central area of the
superficial surface is relatively small. The foramina of the superficial
surface are more abundant than those located on the deep surface,
with an abundance of larger (1-3 mm) and smaller (0.1-0.9 mm)
foramina. A narrow and deep longitudinal crest is present at the deep
surface. This longitudinal crest is laterally bounded by concave sur-
faces. Vascular foramina of the deep surface are less abundant than
those of the superficial face and show scarce large foramina.
The texture of the superficial surface is nearly smooth, showing
only foramina and some grooves. Towards the marginal area, the
texture becomes more irregular, with abundant narrow fibers of
bone, which are almost radially oriented. The texture of the deep
face consists of coarse intercrossing grooves and ridges of irregular
fibers. This pattern changes at the longitudinal crest, where the
grooves and ridges are oriented subparallel to the main axis of the
crest. The fine texture of the deep face consists of fine fibers ori-
ented according to ridges arrangement.
MPCN-PV-804 corresponds to the Morphotype 1 of D’Emic et al.
(2009). This osteoderm is a large and oval-shaped element when
viewed from its superficial face (Fig. 12 E-G). This specimen is
incompletely preserved and its superficial face is badly abraded.
The broken section reveals compact bone without inner large
openings or ducts.
Fig. 11. Posterior dorsal vertebra of an indeterminate Saltasaurini (MPCN-PV-802) in anterior (A, A'), posterior (B, B'), left lateral (C, C'), right lateral (D, D'), and dorsal (E, E') views.
Close-ups of dorsal vertebrae of different saltasaurini from Argentina showing the complex laminae in the lateral side f the vertebra: middle dorsal of Neuquensaurus (F), middle
dorsal of Rocasaurus (G), posterior dorsal of Saltasaurus (H), middle dorsal of MPCN-PV-801 (I), and anterior dorsal of MPCN-PV-802 (J). Light grey parts indicate deep structures.
Dark grey areas indicate broken or missing parts. Scale bars: 10 cm. Abbreviations: Mprsl, Medial prespinal lamina; nc, neural canal; pl, pleurocoel; Pcdl, posterior cen-
trodiapophyseal lamina; Pp-cdf, parapophyseal-centrodiapophyseal fossa; Prpl, prezygoparapophyseal lamina; Pr-cdf, prezygapophyseal-centrodiapophyseal fossa; Pr-ppdf,
prezygapophyseal-parapodiapophyseal fossa; Prsf, prespinal fossa; prz, prezygapophyses; Tprl, intraprezygapophyseal lamina; Upl, unnamed parapophyseal fossa. Numbers
indicate within circles: 1- Posterior centrodiapophyseal lamina; 2- Unnamed parapophyseal lamina; 3- Accessory Posterior centrodiapophsyeal lamina; 4- Prezygapophyseal-
centrodiapophyseal fossa; 5- Parapophyseal-centrodiapophyseal fossa; 6- Postzygapophyseal-centrodiapophyseal fossa and 7-Centroparapophyseal fossa. Red circles shows
laminae and dark circles show fossae. (For interpretation of the references to color in this figure legend, the reader is referred to the Web version of this article.)
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Cretaceous Research 129 (2022) 105029
It is an osteoderm of the kind having a ‘bulb and root’ (e.g., Le Loeuff
et al., 1994; Csiki, 1999; Vidal et al. 2014; Cerda et al., 2015). The
bulb area is very damaged and only preserves a thick marginal
region at the cranial edge. It shows a convex surface with an
ornamentation composed by some small (0.1-0.9 mm) foramina.
The marginal area of the bulb exhibits several grooves and ridges
arranged subparallel to each other and having a superficial-to-deep
orientation, that are continuous along the root. The ridges are
composed of narrow and subparallel fibers of bone. The surface
exhibits a rugose texture at the intersection between the bulb and
root. The deep surface of the root is flat and possesses coarse
intercrossing grooves and ridges of irregular fibers. It shows
numerous foramina of different sizes (0.1-3 mm), and relatively
large vascular foramina (5-10 mm). Three very large vascular
foramina are very close to each other and are placed at the bottom
of the deep surface. This complex of foramina is delimited by inter-
crossing grooves forming a rhomboidal contour.
MPCN-PV-805 represents a Morphotype 1 of D’Emic et al. (2009).
This is ellipsoidal and the largest osteoderm in the sample (Fig. 12
H-J), and is of the kind having a ‘bulb’ morphology (e.g., Salgado and
Coria, 1993; Le Loeuff et al., 1994; Gonza lez Riga, 2003). The su-
perficial face is convex but is badly abraded and does not offer
diagnostic details. The thick marginal region is relatively well-
preserved and it is characterized by having superficial-to-deep
oriented grooves and ridges that are subparallel to each other.
The ridges are composed of narrow fibers of bone. The deep surface
is damaged and lacks anatomical details.
MPCN-PV-806 represents an ellipsoid Morphotype 1 of D’Emic et
al. (2009). This osteoderm is very incompletely preserved and its
deep surface is totally lost (Fig. 12 K-L). The broken sections reveal
compact bone without any large openings or ducts. The osteoderm
presents ‘bulb’ morphology, but a ‘bulb and root’ morphology
cannot be discarded. The superficial face of the osteoderm is
concave in the center of the bulb, and with a radial ornamentation
of foramina (1-3 mm) and fine fibers of bone.
Remarks. Titanosaurs were one of the few sauropods that exhibited
a dermal armor (D’Emic et al. 2009). The presence of osteoderms in
titanosaurs was first reported by Depe ret (1896), who described a
thick osteoderm tentatively assigned to ‘Titanosaurus madagascar-
iensis’ from Madagascar (currently a nomen dubium; Wilson and
Upchurch, 2003). Piveteau (1926) reinterpreted it as belonging to
an ornithischian dinosaur, and the idea of osteoderms in titano-
saurs was abandoned for many years. In this way, the first detailed
description of titanosaur osteoderms from Patagonia was provided
by von Huene (1929), from the locality of Cinco Saltos (Rio Negro
Province, Anacleto Formation) who assigned them to an ankylo-
saurian ornithischian (‘Loricosaurus scutatus’). Definitive evidence
of osteoderms in titanosaurs was presented by Bonaparte and
Powell (1980) for Saltasaurus loricatus from northwestern
Argentina (Bonaparte and Powell, 1980; Powell, 1980, 2003). Since
then, titanosaur osteoderms have been found isolated or associated
with other bones in Africa, Australia, Europe, Pakistan, India,
Madagascar, North and South America (Sanz and Buscalioni, 1987;
Salgado and Coria, 1993; Le Loeuff et al., 1994; Azevedo and Kellner,
1998; Dodson et al., 1998; Csiki, 1999; Gonza lez Riga, 2003;

, F.L. Agnolín et al.
M.A. Rolando, J.A. García Marsa
Fig. 12. Titanosaur osteoderms. MPCN-PV-803 in superficial (A), lateral (B), deep (C) and cranial-caudal (D) views. MPCN-PV-804 in deep (E), lateral (F), and cranial-caudal (G).
MPCN-PV-805 in deep (H), cranial-caudal (I) and superficial (J). MPCN-PV-806 in superficial (K) and lateral (L). Abbreviations. b-r, bulb-root transition. Scale bars: 10 cm.
Malkani, 2003; Powell, 2003; Salgado, 2003; O’Leary et al., 2004;
Gomani, 2005; Marinho and Candeiro, 2005; Salgado et al., 2005b;
Kellner et al., 2006; D’Emic et al., 2009; Curry-Rogers et al., 2011;
Marinho and Iori, 2011; Molnar, 2011; Soto et al., 2012; Díez Díaz et
al., 2013; Lindoso et al., 2013; Vidal et al., 2014; Carrano and D’Emic,
2015; Cerda et al., 2015; Zurriaguz, 2017; Zurriaguz et al. 2017). The
most abundant record of titanosaur osteoderms comes from South
America, especially from Argentina, and several elements have
been collected from the Upper Cretaceous deposits of northern
Patagonia (Cerda et al., 2015). Salgado (2003) described several
isolated osteoderms from localities close to Cerro Matadero (Cinco
Saltos, Lago Pellegrini, Salitral Moreno and Salinas de Trapalco  ; Rio
Negro Province), and more recently, Cerda et al. (2015) realized a
study concerned with the gross morphology, internal micro-
anatomy, and bone histology of titanosaur osteoderms from the
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Cretaceous Research 129 (2022) 105029
Upper Cretaceous of northern Patagonia (Anacleto and Allen
formations).
However, in spite that osteoderms are relatively abundant in the
fossil record of Patagonia, they are often found isolated, and with
only few cases were being closely associated with titanosaur
specimens, and thus, referral of isolated osteoderm to any titano-
saur clade is uncertain. In this regard, Salgado and Coria (1993)
described titanosaurian osteoderms belonging to the genus Aeolo-
saurus, and Salgado et al. (2005a) analyzed osteoderms belonging
to Neuquensaurus. More recently, Zurriaguz et al. (2017) described
seven osteoderms associated to a saltasaurine tail belonging to at
least two morphotypes (keeled and cylindrical).
For this reason, comparisons with previously described osteoderms
clearly referable to a titanosaurian clade and present specimens are
difficult. The specimen MPCN-PV-803 resembles the overall

, F.L. Agnolín et al.
M.A. Rolando, J.A. García Marsa
morphology to those referred to Neuquensaurus australis (Salgado
et al., 2005a; Cerda et al., 2015) and some isolated osteoderms
documented by Cerda et al. (2015), especially MPCA-Pv 6696 and
MPCA-Pv 12342. Nonetheless, MPCN-PV-803 shows a higher
number of large foramina on its superficial face, which is unknown
in other osteoderms from Patagonia.
MPCN-PV-804 is similar in most anatomical details, but smaller, to
the osteoderms described by Le Loeuff et al. (1994) from the Upper
Cretaceous of France and referred to Ampelosaurus, and to the
specimens MPCA-Pv 367/2 and MPCA-Pv 67 described by Salgado
(2003) and Cerda et al. (2015) from Salitral Moreno site (Río
Negro province, Patagonia, Argentina) referred as indeterminate
titanosaurs. MPCA-Pv 367/2 (Salgado, 2003) is a very massive
element, and its “bulb” is comparable in size and shape with that of
MPCN-PV-805 here described.
MPCN-PV-806 is an incomplete element, but resembles on its
slightly concave superficial surface titanosaur osteoderms recov-
ered from Spain (Vidal el al., 2014). However, the transition be-
tween superficial and marginal region is not as abrupt as in the
material described by Vidal et al. (2014). MPCN-PV-806 is notably
different from other known osteoderm previously recovered in
Patagonia.
The possible function of osteoderms in titanosaurs has been much
debated. Some have suggested that osteoderms might have been
related to mineral storage of calcium (Sturkie 1967; Curry-Rogers et
al., 2011; Cerda et al., 2015) which may allow the rapid mobilization
of minerals to the body during the oogenesis (Vidal et al., 2017). In
this regard, paleohistological analyses indicates than the primary
bone tissue of osteoderms is composed of structural fiber bundles
that are ossified by dermal metaplasia and suggests that the
osteoderms were entirely imbedded in the stratum compactum of
the dermis (Cerda et al., 2015). Another highly plausible function
for osteoderms is the defensive role, especially in small species as
saltasaurines or in early ontogenetic stages of larger taxa, which
probably needed extra defense against predators (Marinho and Iori
2011). In this sense, the general consensus is that probably some
titanosaurs were armored forms (Sanz and Buscalioni, 1987; D’Emic
et al., 2009; Curry-Rogers et al., 2011).
In addition, the osteoderm arrangement on titanosaur body is far
from certain. Several different proposals on osteoderm disposi-
tion include osteoderms restricted to the sacro-pelvic region
(Sanz and Buscalioni, 1987) or osteoderms with flat bulbs in the
dorsal and sacral regions and osteoderms with convex bulbs
restricted to the scapular region (Le Loeuff et al., 1994). Salgado
(2003) suggested that symmetric osteoderms formed a sagittal
row and parasagittal rows were composed by asymmetrical
osteoderms. He based this proposal in the fact that in the extant
Alligator mississippiensis, the osteoderms become increasingly
more symmetrical towards the midline of the body. In the case of
keeled osteoderms, because they possessed vaulted margins (in
some cases more ventrally projected than the midline ventral
keel), it is likely that they belonged to the narrower portions of
the body, probably the top of the back, or the middle and distal
regions of the tail (Cerda et al., 2015).
In specimens here reported, it is difficult to indicate to which part of
the body, or to which kind of titanosaur they belong. The keeled
specimen (MPCN-PV-803) based on its shape and morphology may
belongs to the tail (or less probably the midline of the back) of a
saltasaurine titanosaur. The large bulbous specimens (MPCN-PV-
804 and MPCN-PV-805) may belong to the sacro-pelvic area of
Aeolosaurine titanosaurs, based on gross similarities with osteo-
derms reported by Salgado and Coria (1993). The position and af-
finities of osteoderm MPCN-PV-806 are far from certain.
In sum, in spite that the referral of isolated osteoderms to any
titanosaur clade is uncertain, the specimens reported here indicate
21
Cretaceous Research 129 (2022) 105029
that sauropod armors were morphologically diverse at Cerro Mat-
adero, and indirectly indicate the presence of different sauropod
clades.
4. Discussion
The record of Upper Cretaceous sauropods in Neuque n Basin is
the richest from South America. It includes about 32 different
species, most of them belonging to titanosaurs (Salgado and
Bonaparte, 2007; Novas, 2009). To these, is added here the new
genus and species, Menucocelsior arriagadai.
Several previous contributions analyzed the notably diverse
sauropod assemblages from Patagonia (e.g., Bonaparte, 1996;
Leanza et al., 2004; Salgado and Bonaparte, 2007; Novas, 2009;
Juarez-Valieri et al., 2010). Patagonian titanosaurs are almost
grouped in three main clades: the small-sized Saltasaurinae, the
gracile-limbed Aeolosaurini, and the gigantic Colossosauria. In
addition, there are several taxa of uncertain affinities among tita-
nosaurs, including Laplatasaurus, Narambuenatitan, Antarctosaurus,
and others.
At first sight, when the Upper Cretaceous sauropod faunas from
Austral and Golfo San Jorge basins are compared with the Neuque n
basin, some differences come to light (Fig. 13). One of these dif-
ferences is the scarcity of small-sized derived aeolosaurines and
total absence of saltasaurines in the first two basins. At the Neu-
que n Basin gigantic colossosaurians are absent from the record
since the Campanian. Whereas colossosaurians appear to be
abundant and diversified up to the latest Cretaceous at Austral and
Golfo San Jorge basins (see Novas et al., 2019). In spite to that,
relatively large sauropods are not totally extinct from Neuque n
basin, with Antarctosaurus wichmannianus (probably belonging to
the Aeolosaurini-Saltasaurinae lineage; Novas, 2009) and still
innominate taxa (FLA pers. obs.) filling the ecological niche of giant
herbivorous forms during the Maastrichtian. Saltasaurines appear
in the record by the Campanian (Anacleto Formation) but always
retain a low taxonomic diversity.
In the case of Austral Basin, in spite that the record is still not
complete, the sauropods appear to be not very diverse (Fig. 13). For
example, from Chorrillos beds, the single colossosaurian Nullotitan
is known, and represented by several individuals along the strati-
graphic column (Novas et al., 2019), whereas for Cerro Fortaleza
beds only the gigantic and closely-related forms Puertasaurus and
Dreadnougthus were discovered (it should be noted that unpub-
lished remains indicate the presence of an additional smaller tita-
nosaur species) (Novas et al., 2005; Lacovara et al., 2014).
Saltasaurines and aeolosaurines are totally absent from the Austral
basin, and the former appear to be absent south to 42
S latitude
(Salgado and Bonaparte, 2007).
Previous authors noted the extinction of rebbachisaurids at the
CenomanianeTuronian boundary (Coria and Salgado, 2005). At this
boundary, several basal titanosauriforms also became extinct
(Salgado, 2001; Lamanna et al., 2001; Apesteguía, 2002). Posterior
to this extinction, titanosaurian sauropods underwent an extensive
radiation (Salgado et al., 2004; Coria and Salgado, 2005; Gallina and
Apesteguía, 2015; Apesteguía, 2007). In spite that since rebbachi-
saurid extinction titanosaurs increase their diversity, at the Neu-
que n Basin some kind of “evolutionary explosion” occurred at
Campanian times, with aeolosaurines, saltasaurines, and basal
eutitanosaurs being highly diversified. This “explosion” is even
more evident by the latest Cretaceous, were almost all localities
yielded more than one coetaneous sauropod species.
For example, Salitral Moreno (a small and poorly exposed area)
has yielded at least 6 different titanosaurs that roughly coexisted at
the same time (see García and Salgado, 2013), whereas in Salitral
Ojo de Agua, 4 different coeval taxa have been found (Fig. 13). In the

, F.L. Agnolín et al.
M.A. Rolando, J.A. García Marsa Cretaceous Research 129 (2022) 105029

Fig. 13. Figure showing sauropod diversity through time along late Cretaceous beds from each main sedimentary basin from Patagonia. Each sauropod clade is shown separately. At
the bottom, the total number of sauropod species on each sedimentary basin through time is shown. Abbreviations. Aeolos, Aeolosaurini; Ala, Los Alamitos Formation; B.C., Bajada
Colorada Formation; B. de la Carpa, Bajo de la Carpa; Camp, Campanian; Cenom, Cenomanian; Chorr, Chorrillo Formation; Co. Fort, Cerro Fortaleza Formation; Co. Lisandro, Cerro
Lisandro Formation; Colos, Colossosauria; Coni, Coniacian; Maas, Maastrichtian; Rebb, Rebbachisauridae; Salt, Saltasaurini; Sant, Santonian; Titan, non-Saltasaurinae Titanosauria;
Turon, Turonian. The relative ages of the stratigraphical units are a modified and simplified version of previously published information (Garrido, 2010; Novas et al., 2005, 2019;
Casal et al., 2016; Ibiricu et al., 2019). Number of species is based in the case of Austral Basin on Novas et al. (2005, 2019), in the San Jorge Basin on Ibiricu et al. (2011), and the
Neuque n Basin on a large number of publications cited in the main text.

case of Salitral Moreno, there are aeolosaurines, saltasaurines and
four additional titanosaurs of uncertain affinities that do not belong
to the above-mentioned clades. The co-occurrence of large herbi-
vores in coeval beds is striking, and is very different from what is to
be expected if it is employed here extreme actualism (see Bakker,
1977). However, recent work indicates that titanosaurs from
different lineages were rather divergent in body plan and, conse-
quently, niche occupation.
In this sense, saltasaurines, represented in Allen Formation by
Rocasaurus, were very small and robustly built taxa, that had
facultative bipedalism (Powell, 1992, 2003) and possessed an
incredibly pneumatized postcranial skeleton, including both gir-
dles and limb bones (Cerda et al., 2012). Bonatitan reigi that also
comes from Allen Formation, but at the Bajo de Santa Rosa locality
(see Martinelli and Forasiepi, 2004; Salgado et al., 2015), was a
small but gracile-limbed form that lacks the extreme pneumati-
zation observed in saltasaurines, as well as the adaptations for
bipedal stance. Further, its small size, about 3 m in length, in-
dicates that Bonatitan occupied one specific niche not matched by
other titanosaurs.
On the other side, aeolosaurines were medium-sized slender
limbed titanosaurs with a brachiosaur-like ilium and with fore-
limbs longer than hindlimbs (Coria et al., 2013). Further, the tail of
aeolosaurines was strongly modified (Salgado and Coria, 1993) with
very long zygapophyses and anteriorly projected neural arches, that
conferred some rigidity to the tail. Recent analysis indicates that the
tail of aeolosaurines was sigmoidal in contour, presenting a pro-
tonic conformation (Silva-Vidal et al., 2020).
In addition, Antarctosaurus wichmannianus, considered as one of
the largest known dinosaurs described at that time (von Huene,
1929; it probably reached 34 m in length; Mazzetta et al., 2004),
was proportionally slender-limbed and gracile (Powell, 2003). Its
snout was different from other titanosaurs; it was subquadrangular
in shape and show extreme convergences to that of rebbachisaur-
ids, suggesting a highly derived feeding mechanism (Apesteguía,
2004; Martínez et al., 2016).
Thus, in spite of the co-occurrence of different titanosaurs (as
can be seen in Bajo de Santa Rosa, Salitral Moreno and Salitral Ojo
de Agua) their different body plans indicate that they occupied
particular niches and that probably competition for resources was

22

, F.L. Agnolín et al.
M.A. Rolando, J.A. García Marsa
limited, allowing the connivance of several taxa in a single locality
and age. This is not observed in other sites around the world.
The large diversity and morphological disparity of sauropods is
in some way, is also sustained by the high number of different
kinds of osteoderms that have been reported from Maastrichtian
beds at northern Patagonia (Salgado, 2003; Cerda et al., 2015;
Zurriaguz, 2017). However, in the case of osteoderms, their high
morphological diversity at a single site may be explained by
different ways: 1) shape variation within a single individual; 2)
intraspecific variation (e.g., sexual dimorphism), 3) develop-
mental state of the osteoderms (Cerda et al., 2015); and 4)
osteoderm function (e.g. Curry-Rogers et al., 2011; Cerda et al.,
2015). The “developmental state” hypothesis proposed by Cerda
et al. (2015) could be applied to osteoderms recovered from
Salitral Ojo de Agua, especially those included within the Mor-
photype 1 (MPCN-PV-084,805,806; following D'Emic et al., 2009).
However, because these osteoderms were not found in close as-
sociation with any other sauropod bone nor even between them,
this hypothesis should be taken cautiously.
Morphotype 1 resembles bulbous osteoderms widespread
among European titanosaurs (Vidal et al., 2014). This kind of “bulb
and root” osteoderms are probably plesiomorphic for Titanosauria
and for this reason they are widespread in time and space
(Gomani, 2005; Vidal et al., 2014). On the other side, Morphotype
2 osteoderms are present in more derived titanosaurs (i.e. sal-
tasaurines), thus possibly representing a derived condition (Vidal
et al., 2014).
The coexistence of “bulb and roots” and keeled osteoderms
(Morphotypes 1 and 2) is uncommon in the fossil record. Such co-
occurrence was documented for the Maastrichtian Maevarano
Formation, Madagascar, where a basal titanosaur and a saltasaurine
were reported (Curry-Rogers, 2005; Calvo et al., 2007). Salgado
(2003), Cerda et al. (2015), and Zurriaguz (2017) reported for the
Maastrichtian Allen Formation at Salitral Moreno area the simul-
taneous presence of “bulb and root” and “keeled” osteoderms. To
these localities, it is added the Cerro Matadero site which yielded at
least two different titanosaur osteoderm types probably belonging
to both derived and plesiomorphic taxa.
5. Conclusions
The description and analysis of a new sauropod assemblage
coming from the Maastrichtian Allen Formation at Cerro Mataderos
site, close to Salitral Ojo de Agua locality, Río Negro province,
Argentina, resulted in the discovery of unusually high titanosaur
diversity, including the new non-saltasaurine eutitanosaur Menu-
cocelsior arriagadai, the saltasaurine Rocasaurus muniozi, and
indeterminate saltasaurines and aeolosaurines. Further, the co-
occurrence of two different kinds of osteoderms reinforces the
idea of a high sauropod diversity.
It is well known that diversity patterns observed from the fossil
record are influenced by sampling bias. However, the Allen For-
mation localities, particularly Salitral Moreno and Salitral Ojo de
Agua exhibit an unusually high sauropod diversity when compared
with roughly coeval units and sites (e.g., Los Alamitos, Uberaba). As
indicated by García and Salgado (2013), it appears to be certain that
palaeoecological conditions in the area were able to sustain and
promote the flourishing of many species of large-sized herbivores.
Acknowledgements
The authors thank the Arriagada family for guidance to the
fossiliferous outcrops as well as their help during the fieldtrips.
23
Cretaceous Research 129 (2022) 105029
Special thanks to A. Martinelli, M. Ezcurra, L. Salgado, V. Zurriaguz
and M. Militello for their comments on fossil specimens here
described. We also thank M.D. Ezcurra, M.G. Martinelli, M. Reguero,
~ oz for their help during the revision
J. Powell, P. Chafrat and C. Mun
of the paleontological collections under their care. We also thank to
Pablo Chafrat, Franco Migliaro and all the staff of the Museo
Patagonico de Ciencias Naturales (General Roca city) their invalu-
able help during our paleontological expeditions. We sincerely
thank the crew of the LACEV (Laboratorio de Anatomia Comparada
y Evolucion de los Vertebrados), especially to Julia D'Angelo, Ana P.
Moreno, Marcelo P, Isasi, Gonzalo L. Mun ~ oz, Gabriel Lio and Adriel
R. Gentil for their hard work during the field trip. Finally, we thank
to the editor of Cretaceous Research and two anonymous reviewers
for reviewing and considering this publication. This project was
supported by National Geographic Society (Grant Number:
CP050ER17) to MAR and PICT 2018-01390 to FLA.
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